The relationship between dissolved oxygen concentration and maximum size in deep-sea turrid gastropods: an application of quantile regression

Bathymetric gradients in body size are the most well-known patterns of geographic variation in deep-sea organisms. The causes of size-depth relationships remain uncertain, but most have been attributed to rates of nutrient input. Chapelle and Peck (1999, Nature 399:114-115) recently hypothesized that body size in marine invertebrates is a function of dissolved oxygen concentration. We tested this hypothesis by using quantile regression techniques to assess the relationship of dissolved oxygen levels to maximum size in deep-sea turrid gastropods collected from the North Atlantic. Relationships were examined for a group of nine turrid species and within the abundant lower bathyal species Benthomangelia antonia (Dall, 1881). We controlled the analysis for depth because size in deep-sea gastropods varies bathymetrically. When the effects of depth are accounted for statistically, maximum size in B. antonia increases with increasing levels of dissolved oxygen. In turrids as a group, both depth and oxygen appear to explain significant proportions of the variance in maximum size. These findings suggest that a suite of factors, including dissolved oxygen concentration, may influence maximum size in deep-sea organisms.     

Influence of occupant microhabitat on the composition of encrusting communities on gastropod shells

The exploitation of microhabitats is widely considered to increase biodiversity in marine ecosystems. Although intertidal hermit crabs and gastropods may inhabit the same shell type and shore level, their microhabitat may differ depending on the state of the tide. On the south coast of Wales the hermit crab Pagurus bernhardus mainly inhabits the shells of Nucella lapillus (84%). Hermit crab shells had a significantly different encrusting community compared with live N. lapillus shells. At low tide the live gastropods were found on exposed rock surfaces whereas hermit crabs were restricted to tidal pools. Communities encrusting live gastropod shells were characterised by lower species richness and abundance compared with shells inhabited by hermit crabs (12 species found in total). A greater abundance and richness of epibionts was recorded from both shell types during the summer compared with winter. Differences in community composition between shell occupant types were attributed to microhabitats used by gastropods and hermit crabs and the associated desiccation pressures, rather than competitive interactions or shell characteristics. This contradicts earlier studies of subtidal shells where biological processes were considered more important than physical factors in controlling species abundance and richness patterns. The use of rockpool microhabitats by hermit crabs increases the biodiversity of rocky shores, as some species commonly found on hermit-crab-inhabited shells were rare in other local habitats.Communicated by J.P. Thorpe, Port Erin     

Geographical gradients of marine herbivorous fishes: patterns and processes

We present new data and the first rigorous analysis of latitudinal and thermal gradients of diversity, density and biomass of marine herbivorous fishes and review proposed explanatory mechanisms. Consistently, negative relationships between latitude, and positive relationships between sea surface temperature (SST), and relative richness and relative abundance of herbivorous fishes were found worldwide. Significant differences in the strength of gradients of richness and abundance with latitude and SST between tropical and extratropical zones were found consistently across ocean basins. Standardized sampling along the western Atlantic also showed negative relationships between latitude and total density and biomass. The trends, however, are driven by different components of the fish assemblages (i.e., scarids in the Caribbean and acanthurids in Brazil). Patterns of abundance along thermal gradients, generally associated with extensive latitudinal gradients, also were found at the local scale. Feeding rate of the ocean surgeonfish Acanthurus bahianus decreases with temperature more rapidly than the mean metabolic rate of teleost fishes. This relationship suggests a temperature-related physiological constraint. From the new standardized and comparative data presented and the review of the explanatory hypotheses, we conclude that temperature-related feeding and digestive processes are most likely involved in the distribution patterns of herbivorous fishes. Electronic Supplementary Material Supplementary material is available for this article at     

Modelling of concentrations along a moving observer in an inhomogeneous plume. Biological application: model of odour-mediated insect flights

We develop a stochastic model for the time-evolution of scalar concentrations and temporal gradients in concentration experienced by observers moving within inhomogeneous plumes that are dispersing within turbulent flows. In this model, scalar concentrations and their gradients evolve jointly as a Markovian process. Underlying the model formulation is a natural generalisation of Thomson’s well mixed condition [Thomson DJ (1987) J Fluid Mech 180:529–556]. As a consequence model outputs are necessarily compatible with statistical properties of scalars observed in experiment that are used here as model input. We then use the model to examine how insects aloft within the atmospheric boundary-layer can locate odour sources by modulating their flight patterns in response to odour cues. Mechanisms underlying odour-mediated flights have been studied extensively at laboratory-scale but an understanding of these flights over landscape scales is still lacking. Insect flights are simulated by combining the stochastic model with a simple model of insect olfactory response. These simulations show the strong influence of wind speed on the distributions of the times taken by insects to locate the source. In accordance with experimental observations [Baker TC, Vickers NJ (1997) In: Insect pheromone research: new directions, pp 248–264; Mafra-Neto A, Cardé RT (1994) Nature 369:142–144], flight patterns are predicted to become straighter and shorter, and source location is predicted to become more likely as the mean wind speed increases. The most probable arrival time to the source decreases with the mean wind speed. It is shown that scale-free movement patterns arising from olfactory-driven foraging stem directly from the power-law distribution of concentration excursion times above/below a threshold level and are robust with respect to variations in Reynolds number. Flight lengths are well represented by a power law distribution in agreement with the observed patterns of foraging bumblebees [Heinrich B (1979) Oecologia 40(3):235–245].     

Hatching mode and latitude in marine gastropods: revisiting Thorson's paradigm in the southern hemisphere

For much of the last century the developmental mode of marine invertebrates, particularly of prosobranch gastropods, has been thought to reflect a latitudinal pattern known as Thorson's rule; the proportion of species with pelagic larvae decreases with increase in latitude. Although the predictions of this rule have been criticized recently, its validity along latitudinal transects in the world ocean still remains poorly tested. In the present work, we compare the frequency of occurrence of contrasting prosobranch hatching modes (pelagic versus benthic development) along two latitudinal gradients of the subequatorial coastline of South America: the southeast Pacific and the southwest Atlantic. The results are clearly contrasting. While the pattern observed along the Pacific (Chilean) coast fits the predictions of Thorson's rule very well, benthic development predominates all along the Atlantic coast, even at subtropical latitudes. This difference in observed patterns is attributable to the different compositions of the gastropod assemblages on each side of the South American continent, which are determined, in turn, by differences in the ecological conditions on the two sides of the continent. The scarcity of pelagic development among the Atlantic prosobranch gastropods reflects the near-continuous soft-bottom habitat there, and the consequent prevalence of predaceous, soft-bottom taxa that had acquired in their evolution more evolved, non-pelagic patterns of development. In contrast, the Pacific coast is much more heterogeneous, with a diverse mixture of benthic habitats, including rocky substrates that, in part, support grazing taxa. Our results indicate that two factors are important for Thorson's rule to be valid. First, the habitat needs to include rocky substrates, as soft-bottom habitats appear to favour non-pelagic development. Second, a diverse assemblage of taxa need to be compared to avoid the problem of phyletic constraints, which could limit the evolution of different developmental modes. Received: 23 May 2000 / Accepted: 13 October 2000     

Macroinvertebrate diversity of Veli and Kadinamkulam lakes, South Kerala, India

The diversity and distribution pattern of benthic macroinvertebrates in two backwaters viz., Veli and Kadinamkulam of Kerala were assessed using diversity indices. The samples were collected once in three months for a period of two years from six sampling sites (K1, K2, K3, V1, V2 and V3) and community variations were analyzed. Overall, 24 families were identified represented by mollusca, annelida and arthropoda (crustaceans and insects). Among this, dominant taxon was Mytilidae of molluscan family and site-wise dominance was maximum in sites V1 and V2. Richness and abundance were highest in site V2 and lowest in site K2. Diversity index ranged from 0.27 (K2) to 2.33 (V1). The diversity and distribution patterns of certain species were clearly related to water quality as evident from the present study.     

Mechanisms mediating plant distributions across estuarine landscapes in a low-latitude tidal estuary

Understanding of how plant communities are organized and will respond to global changes requires an understanding of how plant species respond to multiple environmental gradients. We examined the mechanisms mediating the distribution patterns of tidal marsh plants along an estuarine gradient in Georgia (USA) using a combination of field transplant experiments and monitoring. Our results could not be fully explained by the "competition-to-stress hypothesis" (the current paradigm explaining plant distributions across estuarine landscapes). This hypothesis states that the upstream limits of plant distributions are determined by competition, and the downstream limits by abiotic stress. We found that competition was generally strong in freshwater and brackish marshes, and that conditions in brackish and salt marshes were stressful to freshwater marsh plants, results consistent with the competition-to-stress hypothesis. Four other aspects of our results, however, were not explained by the competition-to-stress hypothesis. First, several halophytes found the freshwater habitat stressful and performed best (in the absence of competition) in brackish or salt marshes. Second, the upstream distribution of one species was determined by the combination of both abiotic and biotic (competition) factors. Third, marsh productivity (estimated by standing biomass) was a better predictor of relative biotic interaction intensity (RII) than was salinity or flooding, suggesting that productivity is a better indicator of plant stress than salinity or flooding gradients. Fourth, facilitation played a role in mediating the distribution patterns of some plants. Our results illustrate that even apparently simple abiotic gradients can encompass surprisingly complex processes mediating plant distributions.     

Large-scale biodiversity patterns in freshwater phytoplankton

Our planet shows striking gradients in the species richness of plants and animals, from high biodiversity in the tropics to low biodiversity in polar and high-mountain regions. Recently, similar patterns have been described for some groups of microorganisms, but the large-scale biogeographical distribution of freshwater phytoplankton diversity is still largely unknown. We examined the species diversity of freshwater phytoplankton sampled from 540 lakes and reservoirs distributed across the continental United States and found strong latitudinal, longitudinal, and altitudinal gradients in phytoplankton biodiversity, demonstrating that microorganisms can show substantial geographic variation in biodiversity. Detailed analysis using structural equation models indicated that these large-scale biodiversity gradients in freshwater phytoplankton diversity were mainly driven by local environmental factors, although there were residual direct effects of latitude, longitude, and altitude as well. Specifically, we found that phytoplankton species richness was an increasing saturating function of lake chlorophyll a concentration, increased with lake surface area and possibly increased with water temperature, resembling effects of productivity, habitat area, and temperature on diversity patterns commonly observed for macroorganisms. In turn, these local environmental factors varied along latitudinal, longitudinal, and altitudinal gradients. These results imply that changes in land use or climate that affect these local environmental factors are likely to have major impacts on large-scale biodiversity patterns of freshwater phytoplankton.     

Northward shifts of the distributions of Spanish reptiles in association with climate change

It is predicted that climate change will drive extinctions of some reptiles and that the number of these extinctions will depend on whether reptiles are able to change their distribution. Whether the latitudinal distribution of reptiles may change in response to increases in temperature is unknown. We used data on reptile distributions collected during the 20th century to analyze whether changes in the distributions of reptiles in Spain are associated with increases in temperature. We controlled for biases in sampling effort and found a mean, statistically significant, northward shift of the northern extent of reptile distributions of about 15.2 km from 1940-1975 to 1991-2005. The southern extent of the distributions did not change significantly. Thus, our results suggest that the latitudinal distributions of reptiles may be changing in response to climate change.     

Regional differentiation and post-glacial expansion of the Atlantic silverside, <Emphasis Type="Italic">Menidia menidia,</Emphasis> an annual fish with high dispersal potential

The coastal marine environment of the Northwest Atlantic contains strong environmental gradients that create distinct marine biogeographic provinces by limiting dispersal, recruitment, and survival. This region has also been subjected to numerous Pleistocene glacial cycles, resulting in repeated extirpations and recolonizations in northern populations of marine organisms. In this study, we examined patterns of genetic structure and historical demography in the Atlantic silverside, Menidia menidia, an annual marine fish with high dispersal potential but with well-documented patterns of clinal phenotypic adaptation along the environmental gradients of the Northwest Atlantic. Contrary to previous studies indicating genetic homogeneity that should preclude regional adaptation, results demonstrate subtle but significant (F _ST = 0.07; P < 0.0001) genetic structure among three phylogeographic regions that partially correspond with biogeographic provinces, suggesting regional limits to gene flow. Tests for non-equilibrium population dynamics and latitudinal patterns in genetic diversity indicate northward population expansion from a single southern refugium following the last glacial maximum, suggesting that phylogeographic and phenotypic patterns have relatively recent origins. The recovery of phylogeographic structure and the partial correspondence of these regions to recognized biogeographic provinces suggest that the environmental gradients that shape biogeographic patterns in the Northwest Atlantic may also limit gene flow in M. menidia, creating phylogeographic structure and contributing to the creation of latitudinal phenotypic clines in this species.     

湖南省某些稻田土壤固定态铵的剖面分布

利用Silva-Bremner法测定了湖南省某些稻田土壤各剖面的固定态铵量，揭示了其分布规律。固定态铵绝对含量的剖面分布规律有4种类型：第1类为土壤固定态铵随剖面深度增加而增大；第2类是土壤固定态铵随剖面深度增加而减小；第3类是土壤固定态铵随剖面深度变化不明显；第4类是土壤固定态铵在土壤剖面某一土层中明显增大或减少，特别明显的是犁底层的固定态铵量常比耕作层要高。这与成土过程、母质类型以及人为活动等因素有关。固定态铵相对含量分布常从上至下逐渐增加。     

Distribution of a marginal population of <Emphasis Type="Italic">Mytilus edulis</Emphasis>: responses to biotic and abiotic processes at different spatial scales

Physical and biological processes interact to produce pattern in nature. Pattern is scale dependent as processes generating pattern are heterogeneous in time and space. We tested some causes of variation in abundance and distribution of three marginal populations of sublittoral blue mussels, Mytilus edulis, in the non-tidal northeastern Baltic Sea. We studied the role of substrate inclination, perennial algae and siltation along local wave exposure gradients on mussel distribution over a regional salinity gradient. We found marked differences on regional scales (p < 0.001) with lower densities and biomasses of mussels with declining salinity. Along local gradients, mussel densities increased with increasing exposure (p < 0.001) and declining slope and sedimentation (p < 0.01). Site specifically, densities of blue mussels and the perennial red algae, Furcellaria lumbricalis, were positively related, results supported by a colonisation experiment. Also, young post-recruits showed significant relations to adult biomass, wave exposure, algal biomass, bottom slope and sediment cover. Findings showed that the relative importance of the determinants affecting blue mussels at the edge of their range vary with scale and are affected by the density and size structure of mussel populations. The study provides an indication of the types of factors that may be invoked as causes of spatial variation in marginal blue mussel populations and reinforces the need to consider multiple aspects when distributional patterns are assessed.     

Zooplankton biodiversity and lake trophic state: explanations invoking resource abundance and distribution

While empirical studies linking biodiversity to local environmental gradients have emphasized the importance of lake trophic status (related to primary productivity), theoretical studies have implicated resource spatial heterogeneity and resource relative ratios as mechanisms behind these biodiversity patterns. To test the feasibility of these mechanisms in natural aquatic systems, the biodiversity of crustacean zooplankton communities along gradients of total phosphorus (TP) as well as the vertical heterogeneity and relative abundance of their phytoplankton resources were assessed in 18 lakes in Quebec, Canada. Zooplankton community richness was regressed against TP, the spatial distribution of phytoplankton spectral groups, and the relative biomass of spectral groups. Since species richness does not adequately capture ecological function and life history of different taxa, features which are important for mechanistic theories, relationships between zooplankton functional diversity (FD) and resource conditions were examined. Zooplankton species richness showed the previously established tendency to a unimodal relationship with TP, but functional diversity declined linearly over the same gradient. Changes in zooplankton functional diversity could be attributed to changes in both the spatial distribution and type of phytoplankton resource. In the studied lakes, spatial heterogeneity of phytoplankton groups declined with TP, even while biomass of all groups increased. Zooplankton functional diversity was positively related to increased heterogeneity in cyanobacteria spatial distribution. However, a smaller amount of variation in functional diversity was also positively related to the ratio of biomass in diatoms/chrysophytes to cyanobacteria. In all observed relationships, a greater variation of functional diversity than species richness measures was explained by measured factors, suggesting that functional measures of zooplankton communities will benefit ecological research attempting to identify mechanisms behind environmental gradients affecting diversity.     

Functional mechanisms underlying cetacean distribution patterns: hotspots for bottlenose dolphins are linked to foraging

Many studies have shown that the distribution of cetaceans can be closely linked to habitat, but the underlying function of the preferred habitats often remains unclear. Only when behavioural observations are made in relation to habitat types can functional mechanisms behind the habitat use be revealed. Within the range of a bottlenose dolphin (Tursiops truncatus) population off NE Scotland, dolphins show clear preferences for several discrete areas. If the observed patterns of distribution are related to foraging, we predict that behaviour patterns shown by dolphins would reflect this relationship. In this study we identify behaviours of dolphins at the water surface that were related to feeding events, evaluate whether the patterns of distribution were related to foraging and whether they were related the local submarine habitat characteristics. To investigate whether visible surface evidence of foraging behaviour varied spatially, we analysed data collected from 104 regular boat-based surveys made within the Moray Firth, NE Scotland, between 1990 and 2000. To determine whether underlying bathymetry had any influence on the surface behaviour of dolphins, a land-based observation study was carried out in the populations core region of use. The results of this study show that feeding behaviour by dolphins was significantly higher in areas used intensively by dolphins. Furthermore, there were clear relationships between feeding events and the submarine habitat characteristics; certain forms of feeding occur primarily over steep seabed gradients, and in deeper waters during June and July. These results quantitatively support the hypothesis that the distinctive patterns of distribution shown by these dolphins are related to foraging behaviour or opportunities, and that submarine habitat characteristics may be a significant factor in the foraging efficiency of dolphins. Future work should focus on collecting detailed information on the distribution patterns of prey within the study area to allow direct comparisons between predator and prey distributions.Communicated by J.P. Thorpe, Port Erin     

Size-assortative mating and non-reciprocal copulation in a hermaphroditic intertidal limpet: test of the mate availability hypothesis

Mating behaviour and mating patterns are affected to a large extent by body size in both hermaphrodites and gonochorists. Detailed research on mating patterns, mate choice, pre- and post-copulatory sexual selection in marine pulmonates is wanting, thus warranting more attention as a study system in the future. The simultaneous hermaphroditic limpet Siphonaria capensis (Pulmonata: Basommatophora) shows size-dependent fecundity, and acts as a suitable organism to test the effects of body size on mating patterns, mating success, gender expression and reciprocity in hermaphroditic marine gastropods. We mainly used bootstrap resampling techniques to estimate the effects of different factors on mating patterns. In the populations studied, a strong size-assortative mating pattern was observed, where small-scale spatial distribution of potential mating partners (the mate availability hypothesis) could explain 65% of this pattern, while mate choice and mating constraints explaining the rest. No significant difference in mating success between limpets with different body size was found, even though in one population, the sperm recipients were larger than the non-copulating limpets. Interestingly, we found that intromission was non-reciprocal during copulations. This may mean that this species does not conform to the common rule of reciprocity predicted for hermaphrodites, unless there is sex-role alternation between individuals in a mating pair. The mating partners consisted of similar sized, acting males and females, thus without any indication of the body size determining the gender expression. The matings took place in early mornings only at spring tides and the animals were observed laying egg masses only during neap tides. Since siphonariid limpets possess both a spermatheca and a seminal vesicle, which may have either a sperm storing and/or digestive function, post-copulatory sexual selection (e.g., sperm competition) cannot be dismissed.     

Distributions and population structures of two intertidal estuarine polychaetes in the lower St. Lawrence Estuary,with special reference to environmental factors

An intensive study of the spatial distribution ofNereis virens (Sars) andNephtys caeca (Fabricius) was conducted during the spring and autumn of 1986 in the lower St. Lawrence estuary. Statistical analysis showed that spatial variations in density, individual body weight and sexual maturity, particularly in the case ofNereis virens, are correlated with the intertidal level, with certain sediment characteristics, and with the thickness of the colonizable sediment layer. The density ofN. virens increases in an offshore-onshore direction, whereas that ofNephtys caeca decreases in the same direction. For both species, mean body weight increases downshore from the upper intertidal level. Other specific relationships exist in relation to sediment characteristics. Sexually matureNereis virens are found only at the lower intertidal level; sexual maturity inNephtys caeca was not studied. There were no changes in spatial distribution patterns between spring and autumn. The spatial distribution ofNereis virens parallels depth contours and may reflect its ability to inhabit environments which become more physically unstable in an offshore-onshore direction. This spatial distribution is consistent with a model whereby larvae are recruited in the upper intertidal zone and juveniles migrate downshore.     

Sea ice retreat alters the biogeography of the Bering Sea continental shelf.

Seasonal ice cover creates a pool of cold bottom water on the eastern Bering Sea continental shelf each winter. The southern edge of this cold pool, which defines the ecotone between arctic and subarctic communities, has retreated approximately 230 km northward since the early 1980s. Bottom trawl surveys of fish and invertebrates in the southeastern Bering Sea (1982-2006) show a coincident reorganization in community composition by latitude. Survey catches show community-wide northward distribution shifts, and the area formerly covered by the cold pool has seen increases in total biomass, species richness, and average trophic level as subarctic fauna have colonized newly favorable habitats. Warming climate has immediate management implications, as 57% of variability in commercial snow crab (Chionoecetes opilio) catch is explained by winter sea ice extent. Several measures of community distribution and structure show linear relationships with bottom temperature, suggesting warming climate as the primary cause of changing biogeography. However, residual variability in distribution not explained by climate shows a strong temporal trend, suggesting that internal community dynamics also contribute to changing biogeography. Variability among taxa in their response to temperature was not explained by commercial status or life history traits, suggesting that species-specific responses to future warming will be difficult to predict.     

Reproductive isolation among morphotypes of the Atlantic seastar species<Emphasis Type="Italic"> Zoroaster fulgens</Emphasis> (Asteroidea: Echinodermata)

Zoroaster fulgens is a slope-dwelling seastar species that is distributed throughout the Atlantic Ocean. Studies into the population structure and systematics of marine animals have increasingly found that species with a reported cosmopolitan distribution are, in fact, collections of closely related cryptic or sibling species. In the Porcupine Seabight, three morphotypes of Z. fulgens can be found that have a distribution that is stratified by depth. This study investigates the genetic divergence between these morphotypes using sections of the cytochrome oxidase 1 (COI) and 16S regions of the mitochondrial genome. Bathymetrically separated morphotypes of Z. fulgens are reproductively isolated over distances of approximately 1 km while gene flow occurs among morphotypes, along isobaths, over distances of approximately 900 km. Reproductive isolation on the continental slope may have occurred as a result of selection exerted by gradients of depth-correlated physical factors, such as pressure and temperature. However, allopatric speciation with subsequent range expansion may also explain the observed patterns of genetic divergence. Further investigation of radiation within this group may provide important information on the evolution of slope species. Taxonomic revision of the genus is required.Communicated by J.P. Thorpe, Port Erin     

Trait similarity patterns within grass and grasshopper communities: multitrophic community assembly at work

Trait-based community assembly theory suggests that trait variation among co-occurring species is shaped by two main processes: abiotic filtering, important in stressful environments and promoting similarity, and competition, more important in productive environments and promoting dissimilarity. Previous studies have indeed found trait similarity to decline along productivity gradients. However, these studies have always been done on single trophic levels. Here, we investigated how interactions between trophic levels affect trait similarity patterns along environmental gradients. We propose three hypotheses for the main drivers of trait similarity patterns of plants and herbivores along environmental gradients: (1) environmental control of both, (2) bottom-up control of herbivore trait variation, and (3) top-down control of grass trait variation. To test this, we collected data on the community composition and trait variation of grasses (41 species) and grasshoppers (53 species) in 50 plots in a South African savanna. Structural equation models were used to investigate how the range and spacing of within-community functional trait values of both grasses and their insect herbivores (grasshoppers; Acrididae) respond to (1) rainfall and fire frequency gradients and (2) the trait similarity patterns of the other trophic level. The analyses revealed that traits of co-occurring grasses became more similar toward lower rainfall and higher fire frequency (environmental control), while showing little evidence for top-down control. Grasshopper trait range patterns, on the other hand, were mostly directly driven by vegetation structure and grass trait range patterns (bottom-up control), while environmental factors had mostly indirect effects via plant traits. Our study shows the potential to expand trait-based community assembly theory to include trophic interactions.     

Microhabitat-associated variability in survival and growth of subtidal solitary ascidians during the first 21 days after settlement

Newly settled juvenile ascidians were transplanted into shaded and unshaded sites at two subtidal depths to test the hypothesis that photonegative settlement behavior enhances juvenile survival. Silt, filamentous algae and grazing gastropods (Margarites pupillus) were identified as sources of mortality associated primarily with non-cryptic microhabitats. Silt inflicted heavy mortality on all six species tested. Algal overgrowth was important in shallow water, where it increased mortality and depressed growth rates over a 14-d period. Laboratory experiments demonstrated that gastropods can remove juvenile ascidians from the substratum. The sources of post-settlement mortality investigated may help determine distributional patterns of ascidians in the San Juan Islands, Washington, USA, and may also represent selective pressures that maintain negative phototaxis in the behavioral repertoires of ascidian tadpoles.